Effects of climate extremes on the terrestrial carbon cycle: concepts,processes and potential future impacts

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance‐induced mechanisms and processes to also operate in an extreme context. The paucity of well‐defined studies currently renders a quantitative meta‐analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land‐cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground‐based observational case studies reveals that many regions in the (sub‐)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon–climate feedbacks.     

Sexual cannibalism and population viability

Some behaviours that typically increase fitness at the individual level may reduce population persistence, particularly in the face of environmental changes. Sexual cannibalism is an extreme mating behaviour which typically involves a male being devoured by the female immediately before, during or after copulation, and is widespread amongst predatory invertebrates. Although the individual‐level effects of sexual cannibalism are reasonably well understood, very little is known about the population‐level effects. We constructed both a mathematical model and an individual‐based model to predict how sexual cannibalism might affect population growth rate and extinction risk. We found that in the absence of any cannibalism‐derived fecundity benefit, sexual cannibalism is always detrimental to population growth rate and leads to a higher population extinction risk. Increasing the fecundity benefits of sexual cannibalism leads to a consistently higher population growth rate and likely a lower extinction risk. However, even if cannibalism‐derived fecundity benefits are large, very high rates of sexual cannibalism (>70%) can still drive the population to negative growth and potential extinction. Pre‐copulatory cannibalism was particularly damaging for population growth rates and was the main predictor of growth declining below the replacement rate. Surprisingly, post‐copulatory cannibalism had a largely positive effect on population growth rate when fecundity benefits were present. This study is the first to formally estimate the population‐level effects of sexual cannibalism. We highlight the detrimental effect sexual cannibalism may have on population viability if (1) cannibalism rates become high, and/or (2) cannibalism‐derived fecundity benefits become low. Decreased food availability could plausibly both increase the frequency of cannibalism, and reduce the fecundity benefit of cannibalism, suggesting that sexual cannibalism may increase the risk of population collapse in the face of environmental change.     

Minimum viable population sizes and global extinction risk are unrelated

Theoretical and empirical work has shown that once reduced in size and geographical range, species face a considerably elevated risk of extinction. We predict minimum viable population sizes (MVP) for 1198 species based on long-term time-series data and model-averaged population dynamics simulations. The median MVP estimate was 1377 individuals (90% probability of persistence over 100 years) but the overall distribution was wide and strongly positively skewed. Factors commonly cited as correlating with extinction risk failed to predict MVP but were able to predict successfully the probability of World Conservation Union Listing. MVPs were most strongly related to local environmental variation rather than a species' intrinsic ecological and life history attributes. Further, the large variation in MVP across species is unrelated to (or at least dwarfed by) the anthropogenic threats that drive the global biodiversity crisis by causing once-abundant species to decline.     

Scaling the extinction vortex: Body size as a predictor of population dynamics close to extinction events

1. Mutual reinforcement between abiotic and biotic factors can drive small populations into a catastrophic downward spiral to extinction—a process known as the “extinction vortex.” However, empirical studies investigating extinction dynamics in relation to species'' traits have been lacking.
2. We assembled a database of 35 vertebrate populations monitored to extirpation over a period of at least ten years, represented by 32 different species, including 25 birds, five mammals, and two reptiles. We supplemented these population time series with species‐specific mean adult body size to investigate whether this key intrinsic trait affects the dynamics of populations declining toward extinction.
3. We performed three analyses to quantify the effects of adult body size on three characteristics of population dynamics: time to extinction, population growth rate, and residual variability in population growth rate.
4. Our results provide support for the existence of extinction vortex dynamics in extirpated populations. We show that populations typically decline nonlinearly to extinction, while both the rate of population decline and variability in population growth rate increase as extinction is approached. Our results also suggest that smaller‐bodied species are particularly prone to the extinction vortex, with larger increases in rates of population decline and population growth rate variability when compared to larger‐bodied species.
5. Our results reaffirm and extend our understanding of extinction dynamics in real‐life extirpated populations. In particular, we suggest that smaller‐bodied species may be at greater risk of rapid collapse to extinction than larger‐bodied species, and thus, management of smaller‐bodied species should focus on maintaining higher population abundances as a priority.

     

The variability of orchid population size

Orchid population size has long been believed to be highly variable in time and space. Here we present the first quantitative assessment of the variability of orchid population sizes and compare them with plant and animal population data from the literature. Three measures of temporal variation were used (standard deviation of log₁₀ (population size), Hurst exponent and variance in the intrinsic rate of increase). Eight unfertilized plots of Orchis morio in a factorial fertilizer experiment were assessed for variability in population size over a 26 year period. The Hurst exponent across the eight plots gave measures consistent with pink noise well into black noise (0.86 to 1.87). The variation in the intrinsic rate of increase suggested that the expected time to extinction in the plots differed by an order of magnitude. Standard deviation of log₁₀ (population size) across plots was greater than the corresponding temporal variation within plots. Standard deviation of log₁₀ (population size) reached an asymptote between 10 and 20 years. The range of temporal variation between plots in this and other orchid populations was as great as the mean values reported for birds, insects and mammals. These results have highlighted the importance of spatial variation: until we know the relative importance of spatial and temporal variation for more plant species we will be unable to assess whether orchid populations are more variable in size than other species.     

On the interpretation and application of mean times to extinction

As a metric of population viability, conservation biologists routinely predict the mean time to extinction (MTE). Interpretation of MTE depends on the underlying distribution of times to extinction (DTE). Despite claims to the contrary, all information regarding extinction risk can be obtained from this single statistic, the MTE, provided the DTE is exponential. We discuss the proper interpretation of MTE and illustrate how to calculate any population viability statistic when only the MTE is known and the DTE is assumed to be exponential. We also discuss the restrictive assumptions underlying the exponential DTE and the conditions under which alternative models for the DTE are preferable to the conventional (exponential) model. Despite superficial similarities between the exponential and alternative DTEs, several key differences can lead to substantially different interpretations of the MTE.     

种群生存力分析（PVA）的方法与应用

随着人们对资源的加速利用,生境丧失和破碎化导致物种濒危问题日益严重.以岛屿生物地理学为理论起源的种群生存力分析（PVA）,通过分析和模拟种群动态过程并建立灭绝概率与种群数量之间的关系,为濒危物种保护提供了重要的理论依据和研究途径.在过去的几十年中,种群生存力分析已成为保护生物学中一项重要的研究内容.目前种群生存力分析发展稳定,但对其实际预测能力和准确性尚存质疑,应用方面也有待进一步发展.种群生存力分析的进一步完善还需要在理论和方法上的创新,特别是籍于景观生态学和可持续性科学的理念,将空间分析手段、经济社会因素纳入到物种和种群的预测和管理上,从而使其 具有更完整的理论基础和更高的实用价值.为此,本文对种群生存力分析的历史、基本概念、研究方法、模型应用和准确性进行了综述,并提出了有关的研究展望.     

Measurement error and estimates of population extinction risk

It is common to estimate the extinction probability for a vulnerable population using methods that are based on the mean and variance of the long‐term population growth rate. The numerical values of these two parameters are estimated from time series of population censuses. However, the proportion of a population that is registered at each census is typically not constant but will vary among years because of stochastic factors such as weather conditions at the time of sampling. Here, we analyse how such sampling errors influence estimates of extinction risk and find sampling errors to produce two opposite effects. Measurement errors lead to an exaggerated overall variance, but also introduce negative autocorrelations in the time series (which means that estimates of annual growth rates tend to alternate in size). If time series data are treated properly these two effects exactly counter balance. We advocate routinely incorporating a measure of among year correlations in estimating population extinction risk.     

Environmental determinism of year-to-year recruitment variability of European eel Anguilla anguilla in a small coastal catchment, the Frémur River, north-west France

The influence of environmental factors (mainly the river flow) on the year-to-year variability of European eel Anguilla anguilla fluvial recruitment in a small coastal catchment, the Frémur River (north-west France) was examined. A comprehensive survey of catches from fixed traps at two weirs located at 4·5 km (Pont es Omnes Dam) and 6·0 km (Bois Joli Dam) above the river mouth was carried out between 1997 and 2004. Young pigmented elvers (mean ± s.d. total length, 133·7 ± 29·6 mm) were recruited in eel-passes from February to October, but the main runs followed a modal curve from April to September with a peak centred in May to June. Catches varied greatly between years, from 381 to 26 765 elvers. For each trap, a positive linear relationship between monthly mean river flow that preceded the maximal intensity of captures and annual total catches was observed. These relationships explained 73·1% ( P < 0.01) and 89·0% ( P < 0.001) of the year-to-year variability of the recruitment observed in the Pont es Omnes and Bois Joli traps respectively. A significant increase in river flow at the beginning of the migration peak would thus trigger a greater proportion of A. anguilla settled in the estuary and in the downstream zone of the Fremur River to begin their freshwater colonization. The physicochemical roles of changes in river discharge in stimulating upstream migration are discussed. It is concluded that fluvial recruitment in the Fremur River is mainly determined by environmental factors.     

Heterogeneic nature of adult cardiac side population cells

Side population cells have been found in various types of adult tissue including heart and are presumed to be tissue-specific stem/progenitor cells. In the present study, we confirmed the presence of cardiac side population (cSP) cells, which showed both the Hoechst 33342 efflux ability and ABCG2 expression, in adult murine heart. Flow cytometric analysis showed that more than half of cSP cells expressed the endothelial marker VE-cadherin or the smooth muscle markers, α-smooth muscle actin and desmin. In addition, immunohistochemical analysis demonstrated that ABCG2⁺ cells were mainly localized within vascular walls. Quantitative RT-PCR analysis demonstrated that VE-cadherin⁻ cSP cells progressively expressed Nkx2.5 and cardiac troponin T with time in culture. VE-cadherin⁻ cSP cells also expressed mesodermal-mesenchymal-associated markers and differentiated into osteocytes and adipocytes. These results highlight the heterogeneic nature of cSP cells, consisting of vascular endothelial cells, smooth muscle cells, and mesenchymal stem/progenitor cells including potential cardiomyogenic cells.     

The extent of complex population changes in nature

Summary Many models of animal populations show complex yet predictable patterns of density changes under simple and plausible assumptions. Yet one previous attempt to determine the extent and importance of complex dynamics concluded that they were likely only in some laboratory populations, but not in field populations. Ecologists have treated changes more complex than a return to a simple equilibrium, such as the cyclical changes in populations of lynx and voles in the arctic, as special cases. Highly variable populations, such as insects, are usually thought to be driven by unpredictable changes in the weather. Here, we assemble 71 populations counted for over 50 years, and suggest that complex yet predictable population changes are more common than previously thought.     

High variability in patterns of population decline: the importance of local processes in species extinctions

A fundamental goal of conservation science is to improve conservation practice. Understanding species extinction patterns has been a central approach towards this objective. However, uncertainty remains about the extent to which species-level patterns reliably indicate population phenomena at the scale of local sites, where conservation ultimately takes place. Here, we explore the importance of both species- and site-specific components of variation in local population declines following habitat disturbance, and test a suite of hypotheses about their intrinsic and extrinsic drivers. To achieve these goals, we analyse an unusually detailed global dataset for species responses to habitat disturbance, namely primates in timber extraction systems, using cross-classified generalized linear mixed models. We show that while there are consistent differences in the severity of local population decline between species, an equal amount of variation also occurs between sites. The tests of our hypotheses further indicate that a combination of biological traits at the species level, and environmental factors at the site level, can help to explain these patterns. Specifically, primate populations show a more marked decline when the species is characterized by slow reproduction, high ecological requirements, low ecological flexibility and small body size; and when the local environment has had less time for recovery following disturbance. Our results demonstrate that individual species show a highly heterogeneous, yet explicable, pattern of decline. The increased recognition and elucidation of local-scale processes in species declines will improve our ability to conserve biodiversity in the future.     

Harvesting in a resource dependent age structured Leslie type population model

We analyse the effect of harvesting in a resource dependent age structured population model, deriving the conditions for the existence of a stable steady state as a function of fertility coefficients, harvesting mortality and carrying capacity of the resources. Under the effect of proportional harvest, we give a sufficient condition for a population to extinguish, and we show that the magnitude of proportional harvest depends on the resources available to the population. We show that the harvesting yield can be periodic, quasi-periodic or chaotic, depending on the dynamics of the harvested population. For populations with large fertility numbers, small harvesting mortality leads to abrupt extinction, but larger harvesting mortality leads to controlled population numbers by avoiding over consumption of resources. Harvesting can be a strategy in order to stabilise periodic or quasi-periodic oscillations in the number of individuals of a population.     

种群统计随机性和环境随机性对种群绝灭的影响

影响种群绝灭的随机干扰可分为种群统计随机性、环境随机性和随机灾害三大类。在相对稳定的环境条件下和相对较短的时间内,以前两类随机干扰对种群绝灭的影响为生态学家关注的焦点。但是,由于自然种群动态及其影响因子的复杂特征,进一步深入研究随机干扰对种群绝灭的作用在理论上和实践上都必须发展新的技术手段。本文回顾了种群统计随机性与环境随机性的概念起源与发展,系统阐述了其分析方法。归纳了两类随机性在种群绝灭研究中的应用范围、作用方式和特点的异同和区别方法。各类随机作用与种群动态之间关系的理论研究与对种群绝灭机理的实践研究紧密相关。根据理论模型模拟和自然种群实际分析两方面的研究现状,作者提出了进一步深入研究随机作用与种群非线性动态方法的策略。指出了随机干扰影响种群绝灭过程的研究的方向：更多的研究将从单纯的定性分析随机干扰对种群动力学简单性质的作用,转向结合特定的种群非线性动态特征和各类随机力作用特点具体分析绝灭极端动态的成因,以期做出精确的预测。     

Elucidating the impact of temperature variability and extremes on cereal croplands through remote sensing

Remote sensing‐derived wheat crop yield‐climate models were developed to highlight the impact of temperature variation during thermo‐sensitive periods (anthesis and grain‐filling; TSP) of wheat crop development. Specific questions addressed are: can the impact of temperature variation occurring during the TSP on wheat crop yield be detected using remote sensing data and what is the impact? Do crop critical temperature thresholds during TSP exist in real world cropping landscapes? These questions are tested in one of the world's major wheat breadbaskets of Punjab and Haryana, north‐west India. Warming average minimum temperatures during the TSP had a greater negative impact on wheat crop yield than warming maximum temperatures. Warming minimum and maximum temperatures during the TSP explain a greater amount of variation in wheat crop yield than average growing season temperature. In complex real world cereal croplands there was a variable yield response to critical temperature threshold exceedance, specifically a more pronounced negative impact on wheat yield with increased warming events above 35 °C. The negative impact of warming increases with a later start‐of‐season suggesting earlier sowing can reduce wheat crop exposure harmful temperatures. However, even earlier sown wheat experienced temperature‐induced yield losses, which, when viewed in the context of projected warming up to 2100 indicates adaptive responses should focus on increasing wheat tolerance to heat. This study shows it is possible to capture the impacts of temperature variation during the TSP on wheat crop yield in real world cropping landscapes using remote sensing data; this has important implications for monitoring the impact of climate change, variation and heat extremes on wheat croplands.